arXiv subjects

Kirsty Y. Wan

Publications and source records attributed to Kirsty Y. Wan.

13 recordsLinked to original sources

arXiv2026

Tunable asymmetric swimming in biflagellate microswimmers

Many biological microswimmers can modulate their swimming gait to achieve directional control of motility, especially when performing steering towards specific directional cues. This can be achieved without the need for obvious morphological or structural asymmetries in the form of the organism, or in the number or organisation of propulsion-generating appendages such as cilia. In this work, we identify and validate a core principle of asymmetric turning in biflagellate microswimmers: propulsive forces interact constructively to drive translation whilst interacting destructively to drive rotation. We explore the ramifications of this tunable biflagellar swimming mechanism across a range of systems, from a simple, back-of-the-envelope model to a detailed computational representation of an exemplar swimmer. This leads to a markedly general quantitative relation between key drivers of asymmetry, such as ciliary beat frequency, and the curvature of emergent trajectories. We discuss how the model green alga Chlamydomonas reinhardtii, which actuates its two cilia in a symmetric breaststroke for forward swimming, may exploit this feature for phototaxis. Finally, we validate our predictions in a Chlamydomonas-inspired robophysical model, implementing closed-loop control to achieve phototactic turning.

physics.bio-ph↗

arXiv2025

Form and function in biological filaments: A physicist's review

Nature uses elongated shapes and filaments to build stable structures, generate motion, and allow complex geometric interactions. In this Review, we examine the role of biological filaments across different length scales. From the molecular scale, where cytoskeletal filaments provides a robust but dynamic cellular scaffolding, over the scale of cellular appendages like cilia and flagella, to the scale of filamentous microorganisms like cyanobacteria, among the most successful genera on Earth, and even to the scale of elongated animals like worms and snakes, whose motility modes inspire robotic analogues. We highlight the general mechanisms that couple form and function. We discuss physical principles and models, such as classical elasticity and the non-reciprocity of active matter, that can be used to trace unifying themes linking these systems across about nine orders of magnitude in length.

cond-mat.soft↗

arXiv2024

Minimal design of a synthetic cilium

We study a slender filament beating in a viscous fluid with novel curvature-dependent bending stiffness. Our numerical and experimental investigations reveal that such differential stiffness can sustain planar bending waves far along flexible filaments, in stark contrast to the uniform-stiffness case which requires more sophisticated control. In particular, we establish basal actuation as a viable, parsimonious mechanism for generating high-amplitude planar bending waves. Moreover, the resulting beat patterns closely resemble the power-and-recovery strokes of propulsive biological filaments such as cilia, suggesting extensive applications in robotic and engineered systems.

cond-mat.soft↗

arXiv2023

Methods and measures for investigating microscale motility

Motility is an essential factor for an organism's survival and diversification. With the advent of novel single-cell technologies, analytical frameworks and theoretical methods, we can begin to probe the complex lives of microscopic motile organisms and answer the intertwining biological and physical questions of how these diverse lifeforms navigate their surroundings. Herein, we give an overview of different experimental, analytical, and mathematical methods used to study a suite of microscale motility mechanisms across different scales encompassing molecular-, individual- to population-level. We identify transferable techniques, pressing challenges, and future directions in the field. This review can serve as a starting point for researchers who are interested in exploring and quantifying the movements of organisms in the microscale world.

q-bio.CB↗

arXiv2020

Intracellular coupling modulates biflagellar synchrony

Beating flagella exhibit a variety of synchronization modes. This synchrony has long been attributed to hydrodynamic coupling between the flagella. However, recent work with flagellated algae indicates that a mechanism internal to the cell, through the contractile fibres connecting the flagella basal bodies, must be at play to actively modulate flagellar synchrony. Exactly how basal coupling mediates flagellar coordination remains unclear. Here, we examine the role of basal coupling in the synchronization of the model biflagellate \textit{Chlamydomonas reinhardtii} using a series of mathematical models of decreasing level of complexity. We report that basal coupling is sufficient to achieve inphase, antiphase, and bistable synchrony, even in the absence of hydrodynamic coupling and flagellar compliance. These modes can be reached by modulating the activity level of the individual flagella or the strength of the basal coupling. We observe a slip mode when allowing for differential flagellar activity, just as in experiments with live cells. We introduce a dimensionless ratio of flagellar activity to basal coupling that is predictive of the mode of synchrony. This ratio allows us to query biological parameters which are not yet directly measurable experimentally. Our work shows a concrete route for cells to actively control the synchronization of their flagella.

physics.bio-ph↗

arXiv2020

Origins of eukaryotic excitability

All living cells interact dynamically with a constantly changing world. Eukaryotes in particular, evolved radically new ways to sense and react to their environment. These advances enabled new and more complex forms of cellular behavior in eukaryotes, including directional movement, active feeding, mating, or responses to predation. But what are the key events and innovations during eukaryogenesis that made all of this possible? Here we describe the ancestral repertoire of eukaryotic excitability and discuss five major cellular innovations that enabled its evolutionary origin. The innovations include a vastly expanded repertoire of ion channels, endomembranes as intracellular capacitors, a flexible plasma membrane, the emergence of cilia and pseudopodia, and the relocation of chemiosmotic ATP synthesis to mitochondria that liberated the plasma membrane for more complex electrical signaling involved in sensing and reacting. We conjecture that together with an increase in cell size, these new forms of excitability greatly amplified the degrees of freedom associated with cellular responses, allowing eukaryotes to vastly outperform prokaryotes in terms of both speed and accuracy. This comprehensive new perspective on the evolution of excitability enriches our view of eukaryogenesis and emphasizes behaviour and sensing as major contributors to the success of eukaryotes.

physics.bio-ph↗

arXiv2019

Synchrony and symmetry-breaking in active flagellar coordination

Living creatures exhibit a remarkable diversity of locomotion mechanisms, evolving structures specialised for interacting with their environment. In the vast majority of cases, locomotor behaviours such as flying, crawling, and running, are orchestrated by nervous systems. Surprisingly, microorganisms can enact analogous movement gaits for swimming using multiple, fast-moving cellular protrusions called cilia and flagella. Here, I demonstrate intermittency, reversible rhythmogenesis, and gait mechanosensitivity in algal flagella, to reveal the active nature of locomotor patterning. In addition to maintaining free-swimming gaits, I show that the algal flagellar apparatus functions as a central pattern generator which encodes the beating of each flagellum in a network in a distinguishable manner. The latter provides a novel symmetry-breaking mechanism for cell reorientation. These findings imply that the capacity to generate and coordinate complex locomotor patterns does not require neural circuitry but rather the minimal ingredients are present in simple unicellular organisms.

physics.bio-ph↗

arXiv2017

Run stop shock, run shock run: Spontaneous and stimulated gait-switching in a unicellular octoflagellate

In unicellular flagellates, growing evidence suggests control over a complex repertoire of swimming gaits is conferred intracellularly by ultrastructural components, resulting in motion that depends on flagella number and configuration. We report the discovery of a novel, tripartite motility in an octoflagellate alga, comprising a forward gait ($run$), a fast knee-jerk response with dramatic reversals in beat waveform ($shock$), and, remarkably, long quiescent periods ($stop$) within which the flagella quiver. In a reaction graph representation, transition probabilities show that gait switching is only weakly reversible. Shocks occur spontaneously but are also triggered by direct mechanical contact. In this primitive alga, the capability for a millisecond stop-start switch from rest to full speed implicates an early evolution of excitable signal transduction to and from peripheral appendages.

cond-mat.soft↗

arXiv2016

Coordinated Beating of Algal Flagella is Mediated by Basal Coupling

Cilia and flagella often exhibit synchronized behavior; this includes phase locking, as seen in {\it Chlamydomonas}, and metachronal wave formation in the respiratory cilia of higher organisms. Since the observations by Gray and Rothschild of phase synchrony of nearby swimming spermatozoa, it has been a working hypothesis that synchrony arises from hydrodynamic interactions between beating filaments. Recent work on the dynamics of physically separated pairs of flagella isolated from the multicellular alga {\it Volvox} has shown that hydrodynamic coupling alone is sufficient to produce synchrony. However, the situation is more complex in unicellular organisms bearing few flagella. We show that flagella of {\it Chlamydomonas} mutants deficient in filamentary connections between basal bodies display markedly different synchronization from the wild type. We perform micromanipulation on configurations of flagella and conclude that a mechanism, internal to the cell, must provide an additional flagellar coupling. In naturally occurring species with 4, 8, or even 16 flagella, we find diverse symmetries of basal body positioning and of the flagellar apparatus that are coincident with specific gaits of flagellar actuation, suggesting that it is a competition between intracellular coupling and hydrodynamic interactions that ultimately determines the precise form of flagellar coordination in unicellular algae.

cond-mat.soft↗

arXiv2014

Rhythmicity, Recurrence, and Recovery of Flagellar Beating

The eukaryotic flagellum beats with apparently unfailing periodicity, yet responds rapidly to stimuli. Like the human heartbeat, flagellar oscillations are now known to be noisy. Using the alga \textit{C. reinhardtii}, we explore three aspects of nonuniform flagellar beating. We report the existence of rhythmicity, waveform noise peaking at transitions between power and recovery strokes, and fluctuations of interbeat intervals that are correlated and even recurrent, with memory extending to hundreds of beats. These features are altered qualitatively by physiological perturbations. Further, we quantify the recovery of periodic breastroke beating from transient hydrodynamic forcing. These results will help constrain microscopic theories on the origins and regulation of flagellar beating.

cond-mat.soft↗

arXiv2014

Flagellar Synchronization Through Direct Hydrodynamic Interactions

Microscale fluid flows generated by ensembles of beating eukaryotic flagella are crucial to fundamental processes such as development, motility and sensing. Despite significant experimental and theoretical progress, the underlying physical mechanisms behind this striking coordination remain unclear. Here, we present a novel series of experiments in which the flagellar dynamics of two micropipette-held somatic cells of Volvox carteri, with measurably different intrinsic beating frequencies, are studied by high-speed imaging as a function of their mutual separation and orientation. From analysis of beating time series we find that the interflagellar coupling, which is constrained by the lack of chemical and mechanical connections between the cells to be purely hydrodynamical, exhibits a spatial dependence that is consistent with theoretical predictions. At close spacings it produces robust synchrony which can prevail for thousands of flagellar beats, while at increasing separations this synchrony is systematically degraded by stochastic processes. Manipulation of the relative flagellar orientation reveals the existence of both in-phase and antiphase synchronized states, which is consistent with dynamical theories. Through dynamic flagellar tracking with exquisite spatio-temporal precision, we quantify changes in beating waveforms that result from altered coupling configuration and distance of separation. The experimental results of this study prove unequivocally that flagella coupled solely through a fluid medium can achieve robust synchrony despite significant differences in their intrinsic properties.

cond-mat.soft↗

arXiv2013

Lag, lock, sync, slip: the many "phases" of coupled flagella

In a multitude of life's processes, cilia and flagella are found indispensable. Recently, the biflagellated chlorophyte alga Chlamydomonas has become a model organism for the study of ciliary coordination and synchronization. Here, we use high-speed imaging of single pipette-held cells to quantify the rich dynamics exhibited by their flagella. Underlying this variability in behaviour, are biological dissimilarities between the two flagella - termed cis and trans, with respect to a unique eyespot. With emphasis on the wildtype, we use digital tracking with sub-beat-cycle resolution to obtain limit cycles and phases for self-sustained flagellar oscillations. Characterizing the phase-synchrony of a coupled pair, we find that during the canonical swimming breaststroke the cis flagellum is consistently phase-lagged relative to, whilst remaining robustly phase-locked with, the trans flagellum. Transient loss of synchrony, or phase-slippage, may be triggered stochastically, in which the trans flagellum transitions to a second mode of beating with attenuated beat-envelope and increased frequency. Further, exploiting this alga's ability for flagellar regeneration, we mechanically induced removal of one or the other flagellum of the same cell to reveal a striking disparity between the beating of the cis vs trans flagellum, in isolation. This raises further questions regarding the synchronization mechanism of Chlamydomonas.

physics.bio-ph↗

arXiv2013

Antiphase Synchronization in a Flagellar-Dominance Mutant of Chlamydomonas

Groups of beating flagella or cilia often synchronize so that neighboring filaments have identical frequencies and phases. A prime example is provided by the unicellular biflagellate Chlamydomonas reinhardtii, which typically displays synchronous in-phase beating in a low-Reynolds number version of breaststroke swimming. We report here the discovery that ptx1, a flagellar dominance mutant of C. reinhardtii, can exhibit synchronization in precise antiphase, as in the freestyle swimming stroke. Long-duration high-speed imaging shows that ptx1 flagella switch stochastically between in-phase and antiphase states, and that the latter has a distinct waveform and significantly higher frequency, both of which are strikingly similar to those found during phase slips that stochastically interrupt in-phase beating of the wildtype. Possible mechanisms underlying these observations are discussed.

physics.bio-ph↗